Species replacement along a linear coastal habitat: phylogeography and speciation in the red alga Mazzaella laminarioides along the south east pacific

ABSTRACT: BACKGROUND: The Chilean shoreline, a nearly strait line of coast expanding across 35 latitudinal degrees, represents an interesting region to assess historical processes using phylogeographic analyses. Stretching along the temperate section of the East Pacific margin, the region is characterized by intense geologic activity and has experienced drastic geomorphological transformations linked to eustatic and isostatic changes during the Quaternary. In this study, we used two molecular markers to evaluate the existence of phylogeographic discontinuities and detect the genetic footprints of Pleistocene glaciations among Patagonian populations of Mazzaella laminarioides, a low-dispersal benthic intertidal red seaweed that inhabits along ~3,700 km of the Chilean coastal rocky shore. RESULTS: Three main genetic lineages were found within M. laminarioides. They are distributed along the Chilean coast in strict parapatry. The deep divergence among lineages suggests that they could be considered putative genetic sibling species. Unexpectedly, genetic breaks were not strictly concordant with the biogeographic breaks described in the region. A Northern lineage was restricted to a broad transition zone located between 30degreesS and 33degreesS and showed signals of a recent bottleneck. The reduction of population size could be related to warm events linked to El Nino Southern Oscillation, which is known to cause massive seaweed mortality in this region. To the south, we propose that transient habitat discontinuities driven by episodic tectonic uplifting of the shoreline around the Arauco region (37degreesS-38degreesS); one of the most active forearc-basins in the South East Pacific; could be at the origin of the Central/South genetic break. The large beaches, located around 38degreesS, are likely to contribute to the lineages' integrity by limiting present gene flow. Finally, the Southern lineage, occupies an area affected by ice-cover during the last glaciations. Phylogeny suggested it is a derived clade and demographic analyses showed the lineage has a typical signature of postglacial recolonization from a northern glacial refugium area. CONCLUSIONS: Even if environmental adaptation could have strengthened divergence among lineages in M. laminarioides, low dispersal capacity and small population size are sufficient to generate phylogeographic discontinuities determined by genetic drift alone. Interestingly, our results confirm that seaweed population connectivity over large geographic scales does not rely only on dispersal capacity but also seem to depend highly on substratum availability and population density of the receiving locality.     

Microsatellite markers and cytoplasmic sequences reveal contrasting pattern of spatial genetic structure in the red algae species complex Mazzaella laminarioides

Mazzaella laminarioides is a common haploid‐diploid red alga that forms dense beds. This alga has a wide distributional range, covering 3,500 km of the Chilean coast, but is restricted to high rocky intertidal zones. Recently, the existence of three highly divergent genetic lineages was demonstrated for this taxon, and two cytoplasmic markers were used to determine that these lineages are distributed in strict parapatry. Here, using 454 next‐generation sequencing, we developed polymorphic microsatellite loci that cross amplify in all three cytoplasmic lineages. Six sites (i.e., two sites within each lineage) were analyzed using nine microsatellite loci. Our work shows that, although substantial cytoplasmic differentiation occurs within M. laminarioides, the microsatellite loci did not retrieve three nuclear genetic clusters as expected. Indeed, while the northernmost and southernmost cytoplasmic lineages form two strongly divergent nuclear groups characterized by diagnostic alleles, the third cytoplasmic lineage did not form a third nuclear independent group. It is possible that inter‐lineage gene exchange has occurred, particularly at sites along the contact zone between the different cytoplasmic lineages. This nuclear‐cytoplasmic incongruence in M. laminarioides could be explained by incomplete lineage sorting of the nuclear genes or asymmetric introgressive hybridization between the lineages. Finally, highly significant heterozygote deficiencies (suggesting occurrence of intergametophytic selfing) were observed in the three small northernmost sites while the large southernmost sites generally approached panmixia.     

Comparative phylogeography of coastal gobies in the Japanese Archipelago: future perspectives for the study of adaptive divergence and speciation

Ichthyological Research - Phylogeography infers the demographic history of various species by resolving genetic relationships among populations across a geographic range. Comparison of...     

Microbial diversity in coastal sediments during pre- and post-tsunami periods in the south east coast of India

Sediment samples were collected from 12 beaches affected by the 2004 Asian Tsunami in the south-east coast of India between Vanagiri and Nagoor. The objective of the present study is to delineate the microbial diversity in pre- and post-tsunami disaster coastal sediments. The collected marine sediments indicate that the overall microbial diversity is higher in the pre-tsunami sediments. The increase in pathogenic bacteria and fungal species after the tsunami is obscured due to inundation and backwashing of seawater along the coast. The reduction of other microbial diversity after the tsunami is attributed that the coastal and shelf sediments play an important role in the demineralization of organic matter, which supports the growth of microbes. The continuous exchange of ocean water and backwashing of coastal sediments by the tsunami wave probably reduced the pathogenic bacterial diversity in the sediments.     

Comparison of east and west chlorophyll a standing stock and oceanic habitat along the Transition Domain of the North Pacific

Chlorophyll (Chl) a was measured every 10 m from 0 to 150 min the Transition Domain (TD), located between 37 and 45°N,and from 160°E to 160°W, in May and June (Leg 1) andin June and July (Leg 2), 1993–96. Total Chl a standingstocks integrated from 0 to 150 m were mostly within the rangeof 20 and 50 mg m^–2. High standing stocks (>50 mg m^–2)were generally observed westof 180°, with the exceptionof the sporadic high values at the easternmost station. Thetotal Chl a standing stock tended to be higher in the westernTD (160°E–172°30'E) than in the central (175°E–175°W)and eastern (170°W–160°W) TD on Leg 1, but thesame result was not observed on Leg 2. It was likely that largephytoplankton (2–10 and >10 µm fractions) contributedto the high total Chl a standing stock. We suggest that thehigh total Chl a standing stock on Leg 1, in late spring andearly summer, reflects the contribution of the spring bloomin the subarctic region of the northwestern Pacific Ocean. Thedistribution of total Chl a standing stock on Leg 2 was scarcelyaffected by the spring phytoplankton bloom, suggesting thattotal Chl a standing stock is basically nearly uniform in theTD in spring and summer. Moreover, year-to-year variation inthe total Chl a standing stock was observed in the western TDon Leg 1, suggesting that phytoplankton productivity and/orthe timing of the main period of the bloom exhibits interannualvariations.     

中国大陆沿海水鸟栖息地保护优先区及空缺分析

中国大陆沿海湿地是东亚-澳大利西亚候鸟迁徙的重要停歇地,自然湿地丧失与退化已经严重威胁到水鸟生物多样性的稳定性,急需开展物种及其栖息地的保护工作。识别中国大陆沿海水鸟保护优先区,针对保护优先区进行保护优先等级划分,确定目前尚存的保护空缺,是针对性开展水鸟栖息地保护的前提。以中国大陆沿海11省、自治区、直辖市沿岸湿地为研究区,整合鸟类网站、水鸟调查报告和文献中的水鸟调查数据,采用热点分析方法,确定水鸟保护优先区,将每个地块上满足水鸟保护优先区识别标准的物种数量占所有地块中满足标准的物种数量最大值的比例划分成0.5≤P_i≤1、 0.25≤P_i<0.5、0i<0.25三个区间,确定对应的I类、II类和III类水鸟保护优先区保护优先等级;结合国家自然保护地名录,确定水鸟保护优先区保护空缺。结果显示:共有65块中国大陆沿海水鸟保护优先区,记录到满足热点分析标准的水鸟物种共76种,其中受胁物种18种。水鸟保护优先区保护优先等级属于I、II、III类的分别有8块、10块、47块,且水鸟保护优先区保护优先等级属于I...     

海岸线与海岸景观格局对人为干扰度的动态响应——以营口市南部海岸为例

利用多期遥感数据,提取营口南部海岸五期岸线变化信息、分维值,并利用Arcgis10.2中渔网工具创建评价单元。计算景观格局指数,构建人为干扰强度指标,并进一步探讨岸线变化和景观格局变化对人类活动的动态响应。结果表明:(1)研究区围垦导致岸线长度增长、岸线分形维数的增大,1990—2015年四个时段中岸线年增长速率为0.52%、0.53%、4.98%和0.96%;(2)景观格局指数反映2005年之前景观边界、形状复杂程度与破碎程度有所增加;2005年之后景观形状趋于规则化、土地利用趋于均衡化;(3)研究期间强干扰和弱干扰区域面积均有所增加,中等干扰强度区域面积减小;(4)斑块密度、总边缘长度、边缘密度、景观形状指数和平均分维值均与平均干扰强度指数变化同步。Shannon多样性指数与人为干扰强度变化呈反相关;(5)岸线长度变化和岸线分形维数变化都与人类干扰度呈反相关,相关性分别为-0.97和-0.98。     

Diversification in the tropical pacific: comparisons between marine and terrestrial systems and the importance of founder speciation

Patterns of distribution and processes of differentiation haveoften been contrasted between terrestrial and marine biotas.The islands of Oceania offer an excellent setting to explorethis contrast, because the geographic setting for terrestrialand shallow-water, benthic, marine organisms are the same: themyriad islands strewn across the vast Pacific. The size of speciesranges and the geographic distribution of endemism are two biogeographicattributes that are thought to differ markedly between terrestrialand marine biotas in the Pacific. While terrestrial speciesare frequently confined to single islands or archipelagoes throughoutOceania, marine species tend to have wide to very wide distributions,and are rarely restricted to single island groups except forthe most isolated archipelagoes. We explore the conditions underwhich species can reach an island by dispersal and differentiate.Genetic differentiation can occur either through founder speciationor vicariance; these processes are requisite ends of a continuum.We show that founder speciation is most likely when few propagulesenter the dispersal medium and survive well while they travelfar. We argue that conditions favorable to founder speciationare common in marine as well as terrestrial systems, and thatterrestrial-type, archipelagic-level endemism is likely commonin marine taxa. We give examples of marine groups that showarchipelagic level endemism on most Pacific island groups aswell as of terrestrial species that are widespread. Thus boththe patterns and processes of insular diversification are variable,and overlap more between land and sea than previously considered.     

Red-light and blue-light photoreceptors controlling chlorophyll a synthesis in the red alga Porphyra umbilicalis and in the green alga Ulva rigida

Chlorophyll synthesis is stimulated by red light in the green alga Ulva rigida C. Ag. and in the red alga Porphyra umbilicalis (L.) Kützing. Because the effect of red light showed some far-red reversibility in successive red and far-red light treatments, the involvement of phytochrome or a phytochrome-like photoreceptor is suggested. The extent of the response is dependent on exposure and photon fluence rate of red-light pulses. In addition to the effect of red light, a strong stimulation of chlorophyll synthesis by blue light was only observed in Ulva rigida. The effect of blue light shows also some far-red reversibility. In the green alga the accumulated chlorophyll is higher after blue light pulses than after red light pulses. In Porphyra umbilicalis , however, the contrary is observed. In Ulva rigida the involvement of a blue light photoreceptor in addition to phytochrome or a phytochrome-like photoreceptor is proposed. The different responses to red and blue light in both algae are explained in terms of their adaptation to the natural light environment.     

Multiple colonizations, in situ speciation, and volcanism-associated stepping-stone dispersals shaped the phylogeography of the Macaronesian red fescues (Festuca L., Gramineae)

Whereas examples of insular speciation within the endemic-rich Macaronesian hotspot flora have been documented, the phylogeography of recently evolved plants in the region has received little attention. The Macaronesian red fescues constitute a narrow and recent radiation of four closely related diploid species distributed in the Canary Islands (F. agustinii), Madeira (F. jubata), and the Azores (F. francoi and F. petraea), with a single extant relative distributed in mainland southwest Europe (F. rivularis). Bayesian structure and priority consensus tree approaches and population spatial correlations between genetic, geographical, and dispersal distances were used to elucidate the phylogeographical patterns of these grasses. Independent versus related origins and dispersal versus isolation by distance (IBD) hypotheses were tested to explain the genetic differentiation of species and populations, respectively. Genetic structure was found to be geographically distributed among the archipelagos and the islands endemics. The high number of shared AFLP fragments in all four species suggests a recent single origin from a continental Pliocene ancestor. However, the strong allelic structure detected among the Canarian, Madeiran, and Azorean endemics and the significant standardized residual values obtained from structured Bayesian analysis for pairwise related origin hypotheses strongly supported the existence of three independent continental-oceanic colonization events. The Canarian F. agustinii, the Madeiran F. jubata, and the two sister F. francoi and F. petraea Azorean species likely evolved from different continental founders in their respective archipelagos. Despite the short span of time elapsed since colonization, the two sympatric Azorean species probably diverged in situ, following ecological adaptation, from a common ancestor that arrived from the near mainland. Simple dispersal hypotheses explained most of the genetic variation at the species level better than IBD models. The optimal dispersal model for F. agustinii was a bidirectional centripetal stepping-stone colonization pattern, an eastern-to-western volcanism-associated dispersion was favored for F. francoi, whereas for the recently derived F. petraea a counterintuitive direction of colonization (west-to-east) was suggested. The population-based phylogeographical trends deduced from our study could be used as predictive models for other Macaronesian plant endemics with similar distribution areas and dispersal abilities.     

Diets of hooded seals (Cystophora cristata) in coastal waters and drift ice waters along the east coast of Greenland

To provide data on the diets of hooded seals (Cystophora cristata) in the Greenland Sea, seals were collected for scientific purposes on expeditions conducted in the pack ice belt east of Greenland in September/October 1999, 2002 and 2003 (autumn), July/August in 2000 (summer), and February/March in 2001 and 2002 (winter). The results from analyses of stomach and intestinal contents from captured seals revealed that their diet was comprised of relatively few prey taxa. The squid Gonatus fabricii and polar cod (Boreogadus saida) were particularly important, whereas capelin (Mallotus villosus) and sand eels (Ammodytes spp.) occasionally contributed more. These four prey items constituted 60-97% of the diet biomass. Gonatus fabricii was the most important food item in autumn and winter, whereas polar cod dominated the summer diet, with important contributions from G. fabricii and sand eels. The latter was only observed on the hooded seal menu during the summer period, whereas polar cod, which was an important component during the autumn survey, was almost absent from the winter samples. During the latter survey, capelin also contributed to the hooded seal diet. Samples obtained from hooded seals in more coastal waters indicated a more varied diet based on fish such as polar cod, redfish (Sebastes sp.) and Greenland halibut (Reinhardtius hippoglossoides).     

Comparative phylogeography of tenebrionid beetles in the Aegean archipelago: the effect of dispersal ability and habitat preference

Comparative phylogeographical studies in island archipelagos can reveal lineage-specific differential responses to the geological and climatic history. We analysed patterns of genetic diversity in six codistributed lineages of darkling beetles (Tenebrionidae) in the central Aegean archipelago which differ in wing development and habitat preferences. A total of 600 specimens from 30 islands and eight adjacent mainland regions were sequenced for mitochondrial cytochrome oxidase I and nuclear Muscular protein 20. Individual gene genealogies were assessed for the presence of groups that obey an independent coalescent process using a mixed Yule coalescent model. The six focal taxa differed greatly in the number of coalescent groups and depth of lineage subdivision, which was closely mirrored by the degree of geographical structuring. The most severe subdivision at both mitochondrial DNA and nuclear DNA level was found in flightless lineages associated with presumed stable compact-soil habitats (phrygana, maquis), in contrast to sand-obligate lineages inhabiting ephemeral coastal areas that displayed greater homogeneity across the archipelago. A winged lineage, although associated with stable habitats, showed no significant phylogenetic or geographical structuring. Patterns of nucleotide diversity and local genetic differentiation, as measured using Φ_ST and hierarchical amova , were consistent with high levels of ongoing gene flow in the winged taxon; frequent local extinction and island recolonisation for flightless sand-obligate taxa; and very low gene flow and geographical structure largely defined by the palaeogeographical history of the region in flightless compact-soil taxa. These results show that differences in dispersal rate, mediated by habitat persistence, greatly influence the levels of phylogeographical subdivision in lineages that are otherwise subjected to the same geological events and palaeoclimatic changes.     

Glutamate synthase is plastid-encoded in a red alga: implications for the evolution of glutamate synthases

An actively transcribed gene (glsF) encoding for ferredoxin-dependent glutamate synthase (Fd-GOGAT) was found on the plastid genome of the multicellular red alga Antithamnion sp. Fd-GOGAT is not plastid-encoded in chlorophytic plants, demonstrating that red algal plastid genomes encode for additional functions when compared to those known from green chloroplasts. Moreover, our results suggest that the plant Fd-GOGAT has an endosymbiotic origin. The same may not be true for NADPH-dependent GOGAT. In Antithamnion glsF is flanked upstream by cpcBA and downstream by psaC and is transcribed monocistronically. Implications of these results for the evolution of GOGAT enzymes and the plastid genome are discussed.     

Evolution of poecilogony from planktotrophy: cryptic speciation, phylogeography, and larval development in the gastropod genus Alderia

Poecilogony, a rare phenomenon in marine invertebrates, occurs when alternative larval morphs differing in dispersal potential or trophic mode are produced from a single genome. Because both poecilogony and cryptic species are prevalent among sea slugs in the suborder Sacoglossa (Gastropoda: Opisthobranchia), molecular data are needed to confirm cases of variable development and to place them in a phylogenetic context. The nominal species Alderia modesta produces long-lived, feeding larvae throughout the North Atlantic and Pacific, but in California can also produce short-lived larvae that metamorphose without feeding. We collected morphological, developmental, and molecular data for Alderia from 17 sites spanning the eastern and western Pacific and North Atlantic. Estuaries south of Bodega Harbor, California, contained a cryptic species (hereafter Alderia sp.) with variable development, sister to the strictly planktotrophic A. modesta. The smaller Alderia sp. seasonally toggled between planktotrophy and lecithotrophy, with some individuals differing in development but sharing mitochondrial DNA haplotypes. The sibling species overlapped in Tomales Bay, California, but showed no evidence of hybridization; laboratory mating trials suggest postzygotic isolation has arisen. Intra- and interspecific divergence times were estimated using a molecular clock calibrated with geminate sacoglossans. Speciation occurred about 4.1 million years ago during a major marine radiation in the eastern Pacific, when large inland embayments in California may have isolated ancestral populations. Atlantic and Pacific A. modesta diverged about 1.7 million years ago, suggesting trans-Arctic gene flow was interrupted by Pleistocene glaciation. Both Alderia species showed evidence of late Pleistocene population expansion, but the southern Alderia sp. likely experienced a more pronounced bottleneck. Reduced body size may have incurred selection against obligate planktotrophy in Alderia sp. by limiting fecundity in the face of high larval mortality rates in warm months. Alternatively, poecilogony may be an adaptive response to seasonal opening of estuaries, facilitating dispersal by long-lived larvae. An improved understanding of the forces controlling seasonal shifts in development in Alderia sp. may yield insight into the evolutionary forces promoting transitions to nonfeeding larvae.     

Gene lineages and eastern North American palaeodrainage basins: phylogeography and speciation in salamanders of the Eurycea bislineata species complex

Contemporary North American drainage basins are composites of formerly isolated drainages, suggesting that fragmentation and fusion of palaeodrainage systems may have been an important factor generating current patterns of genetic and species diversity in stream-associated organisms. Here, we combine traditional molecular-phylogenetic, multiple-regression, nested clade, and molecular-demographic analyses to investigate the relationship between phylogeographic variation and the hydrogeological history of eastern North American drainage basins in semiaquatic plethodontid salamanders of the Eurycea bislineata species complex. Four hundred forty-two sequences representing 1108 aligned bases from the mitochondrial genome are reported for the five formally recognized species of the E. bislineata complex and three outgroup taxa. Within the in-group, 270 haplotypes are recovered from 144 sampling locations. Geographic patterns of mtDNA-haplotype coalescence identify 13 putatively independent population-level lineages, suggesting that the current taxonomy of the group underestimates species-level diversity. Spatial and temporal patterns of phylogeographic divergence are strongly associated with historical rather than modern drainage connections, indicating that shifts in major drainage patterns played a pivotal role in the allopatric fragmentation of populations and build-up of lineage diversity in these stream-associated salamanders. More generally, our molecular genetic results corroborate geological and faunistic evidence suggesting that palaeodrainage connections altered by glacial advances and headwater erosion occurring between the mid-Miocene and Pleistocene epochs explain regional patterns of biodiversity in eastern North American streams.     

Historical and recent processes shaping the geographic range of a rocky intertidal gastropod: phylogeography,ecology, and habitat availability

Factors shaping the geographic range of a species can be identified when phylogeographic patterns are combined with data on contemporary and historical geographic distribution, range‐wide abundance, habitat/food availability, and through comparisons with codistributed taxa. Here, we evaluate range dynamism and phylogeography of the rocky intertidal gastropod Mexacanthina lugubris lugubris across its geographic range – the Pacific coast of the Baja peninsula and southern California. We sequenced mitochondrial DNA (CO1) from ten populations and compliment these data with museum records, habitat availability and range‐wide field surveys of the distribution and abundance of M. l. lugubris and its primary prey (the barnacle Chthamalus fissus). The geographic range of M. l. lugubris can be characterized by three different events in its history: an old sundering in the mid‐peninsular region of Baja (~ 417,000 years ago) and more recent northern range expansion and southern range contraction. The mid‐peninsular break is shared with many terrestrial and marine species, although M. l. lugubris represents the first mollusc to show it. This common break is often attributed to a hypothesized ancient seaway bisecting the peninsula, but for M. l. lugubris it may result from large habitat gaps in the southern clade. Northern clade populations, particularly near the historical northern limit (prior to the 1970s), have high local abundances and reside in a region with plentiful food and habitat – which makes its northern range conducive to expansion. The observed southern range contraction may result from the opposite scenario, with little food or habitat nearby. Our study highlights the importance of taking an integrative approach to understanding the processes that shape the geographic range of a species via combining range‐wide phylogeography data with temporal geographic distributions and spatial patterns of habitat/food availability.     

The influence of glacial epochs and habitat dependence on the diversity and phylogeography of a coastal dolphin species: <Emphasis Type="Italic">Lagenorhynchus albirostris</Emphasis>

In this paper we use mitochondrial and microsatellite DNA variation to investigate the mechanisms that underlie the evolution of population structure in a highly mobile marine mammal, the white-beaked dolphin. We found moderate genetic diversity (h) at mtDNA, but low nucleotide diversity (π) (0.7320 ± 0.0031 and 0.0056 ± 0.0004, respectively), consistent with expectations for a recent expansion. Analyses based on mismatch distributions further suggested a demographic expansion in the Norwegian-Barents Sea population and a spatial expansion in the British isles-North Sea population, implying distinct demographic histories. F _ST values showed clear differentiation among these two populations, but no difference was found between putative populations separated by the English Channel. Our data suggest a stepwise pattern of expansion, dependent on available coastal habitat. The conservation implications are a need to protect local populations isolated by an expanse of deep water, and in particular, a population along the British coasts and in the North Sea as separate from the North Norway-Barents Sea population. It is also evident that overall diversity was reduced, probably during the last glacial epoch.